Lecture 6

Adaptation
Characteristic that enhances the survival or reproduction of organisms that bear it relative to alternative character states

Adaptations in Action: Examples
Adult
Leaves
Transitional
Leaves
Shingle
Leaves
Philodendron
Two different phenotypes suitable for different conditions at different stages:
• Shingle leaves: capture little light
• Adult leaves: capture more light
Monstera tenuis

Adaptations in Action: Examples
Orchids
Part of the flower is modified to look like a female insect and the flower emits a scent that mimics the attractive sex pheromone of a female wasp
As a male insect “mates” with the flower, pollen is deposited precisely on that part of the insect’s body that will contact the stigma of the next flower visited
Orchid
Chiloglottis formicifera

Adaptations in Action: Examples
Snakes
Most snakes can swallow prey much larger than their heads
The front ends of the two mandibles are not fused (as they are in almost all other vertebrates), but are joined by a stretchable ligament.
Red Diamondback Rattlesnake
Crotalus ruber

Ants
Adaptations in Action: Examples
Weaver ants construct nests of living leaves by the intricately coordinated action of numerous workers
These genetically determined behaviors are adaptations that enhance the reproductive success not of the worker ants that perform them, since the workers do not reproduce, but rather of their mother, the queen, whose offspring include both workers and reproductive daughters and sons
Weaver Ants
Oecophylla

The Nature of Natural Selection
Adaptive biological processes appear to have goals philodendron leaves develop towards a suitable shape and stops developing when the shape is attained, weaver ants act as if they have the goal of constructing a nest
But no conscious anticipation of the future resides in the cell divisions that shape an aroid’s leaf or in the behavior of weaver ants. Rather the apparent goaldirectedness is caused by the operation of a program that controls a process
The information in DNA has been shaped by a historical process of natural selection. Modern biology views the development, physiology and behavior of organisms as the results of purely mechanical processes, resulting from interactions between programmed instructions and environmental conditions or triggers

The Nature of Natural Selection
Natural Selection Any consistent difference in fitness 1 among phenotypically 3 different 4 classes of biological entities 2
1. Fitness Biological entity’s average per capita rate of increase in numbers
The components of fitness generally are: a. Probability of survival to the various reproductive ages b. Average number of offspring produced
Variation in the number of offspring produced as a consequence of competition for mates is often referred to as sexual selection
Fitness is defined for a set of like entities, such as all the individuals with a particular genotype
2. Bio Entities Different kinds of biological entities may vary in fitness, resulting in different levels of selection:
genes/individuals/populations within species/species
3. Phenotypically Requires heritable phenotypes
4. Fitness Diffs Requires variation in fitness

The Nature of Natural Selection
If one neutral allele replaces another in a population by random genetic drift then the bearers of that allele in that population have had a greater rate of increase than the bearers of the other. However, natural selection has not occurred because the genotypes do not differ consistently in fitness: the alternative allele could just as well have increased. There is no average difference between the alleles, no bias toward the increase of one relative to the other
Natural selection resides in the difference in rates of increase among biological entities that is not due to chance. Natural selection is the antithesis of chance!

Experimental Studies of Natural Selection
Natural selection on mutations in the
β-galactosidase gene of Escherichia
coli in laboratory populations maintained in lactose
Mutant with a lower enzyme activity decreased in frequency relative to the wild-type , showing a selective disadvantage
Mutant with higher enzyme activity increased in frequency demonstrating its selective (adaptive) advantage
This illustrates how natural selection is a mindless process without goal.
Adaptation resulted from a difference in the rates of reproduction of different genotypes caused by a phenotypic difference

Experimental Studies of Natural Selection
The courting males of many species of animals have elaborate morphological features and engage in conspicuous displays. Some such features appear to have evolved through female choice of males with conspicuous features, which therefore enjoy higher reproductive success than less elaborate males
Widow Bird
Euplectes progne

Experimental Studies of Natural Selection
Males with shortened tails mate with fewer females than normal males do, and males with elongated tails mate with more females
Natural selection may sometimes lie only in differences in reproductive rate, not survival … sexual selection

Experimental Studies of Natural Selection
“Selfish” Genetic Elements
Are genetic elements that are transmitted at a higher rate than the rest of an individual’s genome and are detrimental (or at least not advantageous) to the organism
SD locus in fruit fly (Drosophila melanogaster)
SD mutant is carried by 95% of the sperm of a heterozygote
In the homozygous condition SD allele is lethal
Natural selection is:
• Differential reproductive success which need not result in adaptation
• Genic selection acts in opposition to individual selection
Development of sperm heads
Drosophila melanogaster

Levels of Selection
Do characters evolve for the survival of the species?
If traits evolve by individual selection then the possibility of future extinction cannot possibly affect the course of evolution
An altruistic trait (a feature that reduces the fitness of an individual that bears it for the benefit of the population species) cannot evolve by group selection
If a population were to consist of altruistic genotypes a selfish mutant (a “cheater”) would increase to fixation, even if a population of such selfish organisms had a higher risk of extinction
The mythical self-sacrificial behavior of lemmings which
(according to popular belief) rush en masse into the sea to prevent overpopulation

Levels of Selection
Traits that benefit the population at a cost to the individual might evolve by group selection?
Pink Selfish Blue Altruistic
The species as a whole might evolve altruism through the greater survival of groups of altruistic individuals even though individual selection within each group would act in the opposite direction

Levels of Selection
Few characteristics have evolved because they benefit the population or species
Individual organisms are much more numerous than the populations into which they are aggregated and they turn over much more rapidly than populations which are formed and become extinct at relatively low rates. Thus the rate of replacement of less fit by more fit individuals is potentially much greater than the rate of replacement of less fit by more fit populations, so individual selection will generally prevail over group selection

Levels of Selection
How do we explain seemingly altruistic behaviors?
One explanation is kin selection. When the altruist enhances the fitness of its relatives, even at some cost to its own fitness, it can increase the frequency of the allele

The Nature of Adaptation
A character is an adaptation if it is a derived character that evolved in response to a specific selective agent
This history-based definition requires that we compare a character’s effect on fitness with those of a specific variant; namely the ancestral character state from which it evolved
One reason for this emphasis on history is that a character state may be a simple consequence of phylogenetic history rather than adaptation. A feature might be beneficial, yet not have evolved for the function it serves today, or for any function at all
P READAPTATION feature that fortuitously serves a new function
Ex. Kea
E XAPTATION preadaptations that have been appropriated to serve a new function
Ex. Wings of guillemots are exaptations for swimming

The Nature of Adaptation
Not all traits are adaptations:
1.
The trait may be necessary consequences of physics and chemistry
2.
The trait may have evolved by random genetic drift rather than natural selection
3.
The trait may have evolved not because it conferred an adaptive advantage but because it was correlated with another feature that did (genetic hitchhiking or pleiotropy)
4.
A character may be the consequence of phylogenetic history

The Nature of Adaptation
One should not assume that a feature is an adaptation unless the evidence favors this interpretation. Several methods are used to infer that a feature is an adaptation for some particular function
1.
Complexity
We often suspect a feature has an adaptive function if it is complex for complexity cannot evolve except by natural selection
Bird’s Eye

The Nature of Adaptation
2.
Design
The function of a character is inferred from its correspondence with the prediction of a model about its function
Cooling in hot tropical climates
Acacia
Banana

The Nature of Adaptation
3.
Experiments
May show that a feature increases fitness relative to individuals in which the feature is modified or absent
4.
Comparative method
Consists of comparing sets of species to pose or test hypotheses on adaptation and other evolutionary phenomena. If a feature evolves independently in many lineages because of similar selective pressure, we can often infer the function of that feature by determining the ecological or other selective factor with which is correlated
Primate Taxa
Sperm competition

What Not Expect of Natural Selection and Adaptation
Some common misconceptions of the theory of adaptive evolution:
1.
The necessity of adaptation
•
An environmental change may set up selection for change in some characteristics
• New adaptations may evolve in an unchanging environment if new mutations arise that are superior to any pre-existing genetic variations
2.
Perfection
Natural selection will not produce absolute perfection. Natural selection cannot fix the best of all conceivable variants if they do not arise and the best possible variants often fall short of perfection due to constraints
3.
Harmony and the balance of nature
Selection at the level of genes and individual organisms is inherently “selfish”. The evolution of a characteristic in a species for the good of another species is not possible
4.
Morality and ethics
Natural selection is not moral or immoral, just or unjust. The naturalistic fallacy (the supposition that what is “natural” Is necessarily “good”, has no legitimate philosophical foundation